lants are chemical machines and one universal characteristic of chemical machines is their sensitivity to

temperature. Temperature, along with light and water,

is one of the most critical factors in the physical environment of plants. This is especially so because plants,
unlike homeothermic animals, are not able to maintain
their tissues at a constant temperature. Environmental
temperature therefore exerts a profound influence on
cellular metabolism and, as a result, plant growth and
their geographic distribution.
All of the chemical machinery of nature—every
individual enzymatic reaction, every metabolic function,
every physiological process—has temperature limits
above and below which it cannot function and an
optimum temperature range where it proceeds at a
maximum rate. Temperature also affects the integrity
of cell structure (especially the structure and properties of membranes), limits the distribution of species in
space and time, and influences the direction of specific
developmental events.
Temperature as an environmental stress and flowering regulator has been discussed in earlier chapters. In
this chapter we will introduce some of the other ways in
which temperature is known to influence plant growth,
development, and distribution. Specific topics include:
• the role of temperature in perennial plants, in particular its role in bud and seed dormancy, and
• some examples of how temperature influences the
geographic distribution of plants.
26.1 TEMPERATURE IN THE
PLANT ENVIRONMENT
Of all the planets, the thermal environment on earth is
particularly fit to give rise to and sustain life. This is
because life functions in an aqueous medium and the
range of temperatures encountered over most of the
earth’s surface generally ensures that sufficient water
is maintained in the liquid state. The temperature
at which biological processes can occur is generally
limited by the freezing point of water on the low
side and the irreversible denaturation of proteins on
the high side. Between these two extremes, a plot of
growth versus temperature for individual organisms
assumes the shape of an asymmetric bell curve, similar
to that for individual enzyme reactions or multiple
enzyme-catalyzed metabolic sequences (Figure 26.1).
In fact, the temperature curve for growth of an
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448 Chapter 26 / Temperature: Plant Development and Distribution
Photosynthesis
Gross
photosynthesis
Net
photosynthesis
Respiration
Growth
Temperature
Tmax
Topt
Tmin
B.
A.
FIGURE 26.1 Temperature and plant growth. (A) A
schematic illustration of the three cardinal temperatures
for plant growth. Typically, the pattern of the growth
curve reflects the pattern of temperature effects on net
photosynthesis. (B) Net photosynthesis is the difference
between gross carbon uptake by photosynthesis and carbon evolution by respiration.
organism effectively represents a composite of the
temperature curves for photosynthesis, respiration, and
other critical metabolic processes. Growth curves thus
exhibit, just as do individual metabolic and enzyme
reactions, the three cardinal temperatures (minimum,
optimum, and maximum). Just as the actual values of
cardinal temperatures vary between different metabolic
processes, the actual values of cardinal temperatures
for growth curves will vary from species to species.
Assuming other factors are not limiting, these cardinal
temperatures generally define the temperature range
over which growth is possible. It is close to the extremes
of this range that plants experience temeprature stress
as described in earlier chapters.
Green plants probably first evolved in the tropical
regions, not so much because of warmer temperatures
(although that may have been a factor), but because the
temperatures there were relatively stable. With time,
plants gradually migrated into the temperate and polar
regions as they adapted to wider variations in temperature on a daily and seasonal basis. Green plants are
now found in regions as extreme as the Antarctic continent and the northern tundra, where temperatures over
much of the year are near or below freezing, and in the
warmest places on earth such as Death Valley (California), where summer temperatures commonly approach
or even exceed 50◦C.
Plants and related organisms may be broadly classified according to their ability to withstand temperature.
Those that grow optimally at lower temperatures
(between 0◦C and 10◦C) are called psychrophiles. The
psychrophiles include primarily algae, fungi, and bacteria. Higher plants generally fall into the category of
mesophiles, whose optimum temperatures lie roughly
between 10◦C and 30◦C. Thermophiles will grow
unhindered at temperatures between 30◦C and 65◦C,
although there are reports of cyanobacteria growing at
temperatures as high as 85◦C. These temperature ranges
apply to hydrated, actively growing organisms. Dehydrated organisms and organs, such as resurrection plants
(Selaginella lepidophylla) and dry seeds with moisture contents as low as 5 percent, are able to withstand a
much
broader range of temperatures for extended periods of
time.
Plants in nature are subjected to a complex mosaic
of fluctuating air and soil temperature regimes such that
it is very difficult to study the effects of temperature in a
natural setting. Air temperature, for example, fluctuates
widely, and often rapidly, depending on the time of
day, cloud cover, season, and other factors. Soil is a
major heat sink as it absorbs and stores solar energy
during the day. At night, some of this heat is radiated
back into the atmosphere, which both cools the soil
and warms the surface. Soil temperature also varies with
the soil structure, organic content, and other physical
characteristics as well as slope and aspect (the direction
it faces with respect to the sun).
Both air and soil temperatures have an impact on
plant growth. Air temperature influences leaf temperature and therefore the rates of photosynthesis, respiration,
and other metabolic reactions. On the other hand,
soil temperature influences germination, root development, and nutrient uptake. For example, maize seeds will
not germinate below about 10◦C and the time required
for germination of winter wheat increases linearly with a
decrease in soil temperature below 25◦C. Several investigators have shown that the uptake of nutrients such
as calcium, boron, nitrogen, and phosphorous increases
with increasing temperature. In cold soils, soybean roots
spread out closer to the soil surface, while i